c 

C49UJ 
ser.2 
no.  7 


tMa  ■:-'■.    WM:M  mM-i  ,   . :'  -,■--■   '■ 


UNIVERSITY  OF  CINCINNATI 


Bulletin  No.  VII 


Publications  of  the  University  of  Cincinnati 

SERIES  II  VOL.  II 

SCIENTIFIC  AND  LITERARY 


QUESTIONS  WITH  ^  AND  Spa  ^ 


BY 

J.    E.    HARRY 


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Q 


AX*** 


Reprinted  from  Studies  in  Honor  of  Basil  Lanneau  Gildersleeve. 

INDICATIVE    QUESTIONS  WITH  ^  AND  apa  rf.1 

The  first  half  of  section  1603  of  Goodwin's  grammar  reads  as 
follows :  "  The  principal  direct  interrogative  particles  are  apa 
and  (chiefly  poetic)  rj.  These  imply  nothing  as  to  the  answer 
expected;  but  apa  ov  implies  an  affirmative  answer  and  apa  py  a 
negative  answer."  The  form  of  these  statements  leads  one  to 
believe  that  apa  pfj  is  a  common  prose  construction,  and,  conse- 
quently, that  the  number  of  examples  of  apa  prj  in  classical 
prose  exceeds  the  number  of  ^'s,  whereas  just  the  reverse  is  true. 
In  section  1015  of  Hadley-Allen  the  sentence  apa  prj  foa/3a\W0at 
tiogas;  is  cited  without  a  hint  as  to  the  extent  of  the  use  of  apa  prj  either 
in  prose  or  poetry.  Kiihner,  587,  11,  speaks  only  of  pi]  and  has 
only  three  words  on  this :  "  erst  seit  Aeschylus,"  though  to  be 
sure,  in  587,  14,  he  says  that  apa  occurs  "erst  in  der  nachhome- 
rischen  Sprache."  All  other  grammars,  both  German  and  English, 
are  as  silent  on  this  subject  as  Kiihner.  The  lexicons  either 
furnish  little  information  or  are  misleading.  (Cf.  Amer.  Journ.  of 
Philol.  Ill,  515  and  XIX,  233.)  Commentaries  show  as  little 
sense  of  proportion  in  respect  to  the  usage  of  these  interrog- 
ative particles  as  the  grammarians. 

Dyer  on  Apology  25  A  remarks  that  "  questions  with  ^  take  a 
neg.  answer  for  granted,"  and  on  Crito  44  E  "  apa  pfj  looks  for  a 
neg.  answer,  but  it  may  also  convey  an  insinuation  that  in  spite  of 
the  expected  denial  the  facts  really  would  justify  an  affirmative 
answer."  There  is  no  intimation  of  the  limitations  of  both  prj  and 
apa  prj.  On  apa  firj  in  Memorabilia  11,6,  34  Winans  has  nothing  to 
say;  on  IV,2, 10  he  refersto  the  grammars  of  Goodwin  and  Hadley; 
and  on  pfj  in  IV,  2,  12  to  Goodwin's  Moods  and  Tenses  46  n.  4, 

xMy  attention  was  first  directed  to  this  subject  by  Dr.  C.  W.  E.  Miller, 
who  pointed  out  to  me  the  rare  use  of  juy  (apa  pjj)  as  an  interrogative 
particle  in  Classical  Greek,  and  told  me  that  as  the  result  of  observations 
in  this  direction  he  felt  certain  that,  with  the  exception  of  perhaps  a  solitary 
example  in  Demosthenes,  the  construction  was  not  found  in  the  Attic 
orators,  and  that  Plato  was  about  the  only  prose  writer  that  employed  it  to 
any  noteworthy  extent. — J.  E.  H. 


-O 


^&^ 


428  /.  E.  HARRY. 

where,  he  remarks,  "  another  interpretation  is  given,  however, 
reading  dvvapai  with  Kiihn.  and  several  MSS."  But  Goodwin 
reads  Svvapai  in  the  edition  of  1890  (268). 

With  the  exception  of  three  examples  in  Xenophon,  apa  $  does 
not  occur  in  prose  outside  of  Plato ;  and  in  the  2442  pages  of 
the  extant  works  of  this  author  (Teubner  text)  only  ten  examples 
of  the  construction  are  found,  two  of  these  being  in  spurious 
dialogues  (Anterastae  and  De  Virtute).  The  Phaedo  contains 
three  of  the  remainder ;  two  of  these  may  be  counted  as  one — 
64  C,  where  apa  prj  aXXo  ti  fj  is  used  and  then  repeated  in  toto  in 
resuming  the  question ;  the  third  is  found  in  103  C.  The  remain- 
ing five  are  distributed  as  follows:  Crito  44  E,  Parmenides  163  C, 
Charmides  174  A,  Lysis  213  D,  Republic  405  A.  The  indicative 
is  used  in  all  the  examples  except  the  second  one  of  Phaedo  64  C, 
which  has  the  subjunctive,  like  the  examples  of  simple  ntf  in 
cautious  questions.1 

The  frequency  of  occurrence  of  the  interrogative  particle  apa, 
alone  and  combined  with  ov,  ye,  ovv  and  prj,  in  the  dialogues  of 
Plato  may  be  seen  from  the  following  conspectus  : 


apa 

apa.  ye 

ap    ovv 

T      )         > 

ap    ov 

Total. 

apa 

Euthyphro2 

7 

I 

3 

4 

14 

.    . 

Apology 

2 

.   • 

1 

.  . 

3 

.   • 

Crito    . 

3 

I 

1 

1 

6 

I 

Phaedo 

*3 

.   . 

9 

13 

33 

3 

Cratylus 

11 

2 

12 

19 

4i 

Theaetetus 

25 

.    . 

11 

11 

44 

Sophistes .    . 

15 

.    . 

6 

29 

48 

Politicus  . 

13 

I 

8 

17 

38 

Parmenides . 

13 

2 

20 

26 

55 

1 

Philebus  . 

18 

.    . 

21 

28 

65 

Symposium  . 

5 

I 

3 

.  • 

9 

Phaedrus . 

1 

I 

6 

9 

13 

Alcibiades  I     . 

17 

.    . 

13 

6 

34 

Alcibiades  II  . 

2 

3 

7 

5 

16 

1  Goodwin  (M  T  268)  and  Weber  cite  all  the  examples  except  Craty 
429  C  pi)  yap  ovde  tovto  ah  y,  to  tovtov  (f>dvat  'Ep/^oye^^  elvai,  el  /xi)  hanv ; 

2ap'  ovv  ov  (14  D)  is  counted  twice.     Hence  the  apparent  mistake  in 
total  column.     So  also  in  Leges  and  De  Virtute. 


INDICATIVE  QUESTIONS   WITH  m  AND  apa  ^     429 


apa          apa  ye      ap    ovv 

op    ov 

Total. 

Hipparchus  . 

4 

2 

.  . 

6 

Anterastae 

1 

3 

I 

5 

Theages    .    . 

2 

2          .  . 

4 

8 

Charmides 

5 

1             3 

2 

11 

Laches      .    . 

4 

.  . 

I 

5 

Lysis     . 

9 

1           10 

2 

21 

Euthydemus 

13 

2           13 

2 

30 

Protagoras    . 

n 

7 

8 

26 

Gorgias     . 

27 

22 

I 

50 

Meno     . 

11 

5 

6 

21 

HippiasMaior 

4 

8 

6 

16 

Hippias  Minor 

2 

.  .             2 

2 

5 

Ion     . 

2 

.  . 

.  . 

2 

Republic 

42 

4          95 

81 

190 

Timaeus 

1 

3 

3 

6 

Minos 

1 

.  .            2 

2 

5 

Leges  . 

29 

4           33 

59 

113 

Epinomis 

1 

.  . 

3 

4 

Epistolae 

2 

1             1 

.  . 

4 

De  Iustitia 

1 

1           •  • 

6 

8 

De  Virtute  . 

3 

3 

1 

6 

Demodocus 

1 

.  .             1 

.  , 

2 

Sisyphus 

3 

3            3 

2 

11 

Alcyon 

.  . 

1           .  . 

.  . 

1 

Eryxias 

10 

4            5 

19 

apa  prj 


334 


36 


342 


360 


994 


10 


It  will  be  seen  from  the  table  that  apa,  ap*  ovv  and  ap'  ov  nearly 
balance  each  other ;  there  are  nearly  twice  as  many  examples  of 
apa  as  of  ap'  ov,  and  only  one  out  of  every  hundred  of  the  apa's  is 
followed  by  pq. 

There  are  104  examples  of  apa  in  the  orators  (including  both 
genuine  and  spurious  speeches).  Of  these  22  are  followed  by  ov. 
*Apd  ye  appears  17  times  and  ap'  ovv  23.  Demosthenes  has  a 
greater  number  of  Spa's  than  all  the  others  together  (64) ;  half  of 
them  are  found  in  orations  XVIII-XXIV ;  and  ten  are  followed 
by  ov.  There  is  little  variation  in  the  figures  for  the  rest  of  the 
orators  (except  Antiphon,  in  whom  the  particle  does  not  occur), 


43°  /.  E.  HARRY. 

Andocides  having  two  examples  (both  without  ov),  Lysias  seven 
(one  negative),  Isocrates  five  (one  neg.),  Isaeus  five  (one  neg.), 
Lycurgus  six  (three  negatives),  Aeschines  five  (one  neg.), 
Hyperides  four  (all  neg.),  Dinarchus  four  (no  negatives). 

In  the  historians  apa  hardly  makes  its  appearance — twice  in 
Herodotus  (apa  III,  50 ;  apJ  ov  IX,  27)  and  only  once  in  Thucyd- 
ides  (I,  75,  1,  where  apa  =  ap  ov,  as  in  Sophocles,  O.  C.  753,  780, 
Aristophanes,  Birds  797). 

Xenophon  has  90  examples  of  apa  [36  of  simple  apa,  15  of  ap  ov, 
2  of  apa  firj,  26  of  ap  ovv  (including  one  apy  ovv  ...  firf)f  and  1 1  of  apa 
ye].  More  than  half  of  these  (48)  occur  in  the  Memorabilia.  The 
rest  appear  as  follows:  Anab.  4,  Cyropaed.  18,  Hellen.  1,  minor 
works  19.  Of  the  15  examples  of  ap  ov,  eight  belong  to  the 
Memorabilia,  three  to  the  Anabasis,  one  to  the  Cyropaedia  and 
three  to  the  minor  works.  Ten  examples  of  the  combination  ap'  av 
(followed  by  the  optative)  are  found  in  the  Cyropaedia  alone. 
The  references  for  the  three  instances  of  apa  pr]  are  Mem.  II,  6, 
34;  IV,  2,  10;  and  Anab.  VII,  6,  5. 

Interrogative  ^  occurs  neither  in  the  orators  1  nor  in  the  his- 
torians. Even  paw,  which  is  commoner  in  Plato  than  pi)  and  must 
be  regarded  as  differing  from  pi)  ovv  (pwv  prj,  p5>v  ov  and  p&v  ovv  are 
not  rare),  does  not  appear  in  the  orators,  historians  or  Xenophon. 

There  are  twenty-four  examples  of  \u\  interrogative  in  Plato. 
Of  these  the  greatest  number  is  in  the  Republic  (6);  the  Protag- 
oras comes  next  with  five ;  two  each  are  found  in  Euthydemus, 
Gorgias,  Meno,  and  Apology ;  one  each  in  Phaedo  and  Hippias 
Major,  and  three  in  the  Theaetetus  (not  counting  the  repetition 
in  146  E).  In  Meno  89  C  (pr)  tovto  ov  ko\5>?  oopoXoyrjaapevy)  ov  and 
Ka\S>s  coalesce,  as  does  ov  and  roiavrrjv  in   Protagoras  312  A  firj  ov 

roiavrrjv    v7ro\ap(3dv€is    o~ov    ttjv     p-dBrjo-tv     eaeaOai )     (which,    however, 

Goodwin  considers  declarative).  Over  against  these  24  examples 
of  pr)  there  are  83  instances  of  /*£?,  which  include  28  occurrences 
of  pcov  ov,  5  of  pS>v  prj,  and  18  of  fiwv  ovv,  this  last  embracing  8 
instances  of  pS>v  ovv  ov. 

1  There  is  an  example  of  firj  with  the  past  indie,  in  Dem.  XX,  160  (rl;  p.Tj 
ml  ra  peXkovr'  j>deig  ;),  but  the  passage  is  possibly  corrupt.  The  form  of  the 
rhetorical  vrroipopa  immediately  following  indicates  that  the  preceding 
question  was  not  put  as  it  appears  in  our  MSS.  Many  readings  suggest 
themselves,  e.  g.  rl  del  ml  to,  tikXkovr''  rjdrj  ; 


INDICATIVE  QUESTIONS  WITH  M  AND  apa  pij.     43 1 

All  the  questions  introduced  by  prj  in  Xenophon  occur  in 
Memorabilia  IV,  2,  10  (except  one  in  III,  11,  4  prj  xeiporexvai  raw;), 
and  the  four  found  here  are  merely  a  continuation  of  Socrates' 

question  *Apa  pfj  larpos   (SC.   (3ov\ei,  Or   emOvpels,  yeveaOai)  ; 1      pap,  as 

has  been  stated  above,  does  not  occur  in  Xenophon. 

So  much  for  the  classical  prose  writers.  Let  us  now  direct 
our  attention  to  the  poets.  Though  apa  prj  is,  as  we  have  seen, 
rare  in  prose,  it  is  still  rarer  in  poetry,  there  being  only  three 
examples  in  the  whole  range  of  epic,  lyric  and  dramatic 
literature.  No  instance  can  be  cited  from  Homer ;  none  from  the 
melic  poets;  none  from  Aristophanes.  It  appears  twice  in 
Sophocles  (El.  446,  Ant.  632)  and  once  in  Aeschylus  (Septem 
208).  As  for  ap  ov,  Aeschylus  has  not  a  single  example,  Sophocles 
but  three,  Euripides  five,  whereas  Plato  has  360.  Simple  apa 
occurs  ten  times  in  Aeschylus ;  there  are  38  examples  in  Sophocles 
and  52  in  Euripides— just  100  in  all.  There  are  48  apa's  (seven 
of  these  followed  by  ov)  in  Aristophanes,  but,  as  has  been  stated, 
not  a  single  apa  pi]. 

Simple  p?  (without  a  preceding  interrogative  particle)  is  not 
found  before  Aeschylus,  and  in  all  the  tragic  poets  occurs  but 
six  times,  four  of  these  being  in  Aeschylus  (P.  V.  247,  959,  Pers. 
344,  Suppl.  295),  one  in  Sophocles  (Trach.  316), 2  one  in  Euripides 
(Hipp.  799).  In  Aristophanes  there  is  but  one  example,  and  that 
is  found   in  the  brogue  of  the   Scythian  archer  (Thesm.  11 14 

(TKe^rai    ro  kvo-to'  pr)   n   piKrbv    rraivcrai ;).       The    compound    pa>v,    On 

the  other  hand,  (used  only  by  the  Attic  writers),  can  not  be 
classed  with  pi),  for,  although  it  is  not  employed  by  any  prose 
writer  except  Plato,  the  particle  occurs  frequently  in  comedy  (27 
examples  in  Aristophanes)  as  well  as  in  tragedy  (41  examples). 
The  fact  that  p5>v  occurs  33  times  in  Euripides  and  only  five  and 
three  times  in  Sophocles  and  Aeschylus  respectively  (together 
with  its  use  in  comedy  and  its  absence  from  the  orators  and  histo- 
rians) seems  to  indicate  that  it  belongs  to  the  sermo  familiaris. 

xThere  is  another  example  in  the  Oeconomicus  (XII,  1  prj  ae  Karaaulvo) 
anievai  rjdr)  /3ov2,6pevov;).  This  may,  however,  be  taken  as  a  hortatory  sub- 
junctive, and  so  Holden  explains  (although  in  his  text  the  sentence  is 
interrogative),  translating  "let  me  not  detain  you,"  and  referring  to  Goodwin 
253  (1344).  Kiihner  and  Dindorf  regard  the  sentence  as  a  question.  In 
Mem.  IV,  2,  12  pi)  ovv  .  .  .  ov  dvvupat  /ere.  the  mood  is  the  subjunctive. 

2  The  verb  in  this  passage  is  unexpressed. 


432  /•  E.  HARRY. 

Mav  ovv  is  found  twice  in  Aeschylus  and  once  in  Euripides ;  pa>v  ov 
occurs  but  twice  in  the  tragic  poets  (Eur.  Med.  733,  Troad.  714), 
Ia£>v  fir]  not  at  all. 

The  interrogative  7  occurs,  of  course,  much  more  frequently 
in  the  tragic  poets  than  in  prose  (25  times  in  Aeschylus,  61  times 
in  Sophocles,  and  74  times  in  Euripides).  Aristophanes  again 
comes  near  the  prose  norm  with  hardly  a  dozen  examples. 

If  I  can  trust  to  a  rapid  reading  of  Aristotle,  neither  ^  nor  apa 
fxrj  appears  in  his  writings.  The  same  may  be  said  of  Callimachus, 
Apollonius  Rhodius,  Lycophron,  Theocritus,  Bion,  Moschus, 
Polybius,  and  Diodorus  Siculus  (2043  Teubner  pages).1 

In  Theophrastus  apa  prj  does  not  occur  at  all  and  prj  is  found 
but  once,  and  that  in  one  of  the  Characters  (nept  Aoyonoiias),  where 
the  author  is  giving  a  sample  of  ordinary  small  talk,  and  puts 
in  the  mouth  of  his  character  the  words  prj  Xeycrai  Kaivorepov',  Im- 
mediately thereafter  Foss  would  read  prj  dyaOd  ye  eVn  to.  Xeyopeva ; 
but  the  MSS  have  Ka\  pfjv  instead  of  prj. 

Yet  in  spite  of  the  fact  that  in  the  whole  domain  of  Greek  liter- 
ature, from  Homer  down  to  the  time  of  Christ,  a  period  of  one 
thousand  years,  apa  prj  appears  but  three  times  in  poetry  and 
11  (13)  times  in  prose,  a  celebrated  scholar  (Blaydes)  desires 
to  emend  a  perfectly  intelligible  sentence  in  Sophocles  apd  pov 

pepvrjaOe;  (O.  T.  I401)  SO  as  tO  read  apa  pr)  pepvrjcrOe; 

When  we  come  to  the  New  Testament  we  have  a  different  story 
to  tell :  pi)  in  questions  is  common — eight  times  in  Matthew,  four 
in  Mark,  six  in  Luke,  twenty-one  in  John,  four  in  James,  eight  in 
Romans,  fourteen  in  I  Corinthians  and  four  in  II  Corinthians. 
All  of  these  are  with  the  indicative.  The  sum  total,  then,  of  ques- 
tions with  prj  in  the  New  Testament  is  sixty-nine,  a  greater  number 
than  in  all  the  prose  and  poetry  of  the  ten  centuries  preceding. 

All  the  examples  of  m  in  the  New  Testament  are  found  in  eight 
books,  the  four  gospels  containing  more  than  half  of  the  whole 
number  (39).  About  one-third  of  the  number  (21)  are  in  John 
alone.     In  about  one-half  of  the  cases  (32)  the  verb  is  one  of  the 

xNot  unlike  the  behavior  of  apa  pij  is  that  of  aXko  tl  and  aXko  n  fj.  These 
phrases  do  not  appear  to  any  extent  outside  of  Plato.  There  is  not  a  single 
example  in  the  orators  except  Lysias  (two  instances  only,  one  of  these  in 
a  genuine  speech  and  supporting  the  thesis  that  the  phrase  belongs  to  the 
language  of  everyday  life,  the  other  in  a  spurious  speech)  and  the  un- 
rhetorical  orator,  Andocides. 


INDICATIVE  QUESTIONS   WITH  m  AND  apa  pi).      433 

three  that  are  most  common  in  the  speech  of  everyday  life  (be, 
can,  have).  An  even  dozen  of  the  p^s  appear  in  the  form  oi  pi)  ™. 
The  double  negative  pi)  ov  is  found  in  Romans  x,  18.  The  nega- 
tive ovxi  is  very  frequently  the  introductory  word  of  a  sentence; 
and  apa  ye   is  found  in  Acts  viii,  30. 

The  behavior  of  the  particles  in  later  Greek  is  similar  to  their 
conduct  in  the  pre-Christian  period.  Dionysius  of  Halicarnassus, 
who  sought  to  revive  a  true  standard  of  Attic  prose,  has  not  a 
single  example  of  either.  In  Plutarch  (3670  pages  in  the  Teubner 
text)  apa  prj  does  not  occur  (though  apa  alone  does),  pi)  only  once, 
Alexander  XXVIII  pr)  n  o-v  toiovtov  6  tov  Ai6s; 

In  the  sophist  Dio  Chrysostomus  we  find  two  examples  of  apa  pi) 

and  six  of  pi)  :  XXXII  (683  R)  apa  ye  pr)  AaKedaipoviovs  pipiiaOe  \  LVII 
(296  R)  apa  pi)  d\a£6va  TiziroirjKe.  tov  NeVropa ;  IX  (294  R)  pr)  ovv 
davpd£ovo~iv  avTov)  X  (306  R)  pr)  yap  eKeivos  eXvcre  to  alviypa)  XIV 
(438  R)  pr)  ovv  o~v  (firjs  eXevdepov  eivai  tov  av8pa  tovtov  )  XXX  (54^  ^) 
aXXci  pi)  ti  vpas  Au7rei ;  XXXII  (676  R)  pr)  to.  a>Ta  €7raXr)\iTTTai  twv 
exet  \    LVIII  (301  R)  M  °vp  clvtos  ye  alpel ', 

Even  Lucian,  in  spite  of  the  fact  that  he  wrote  the  best  Attic 
prose  that  had  been  written  for  four  hundred  years,  is  not  fault- 
less. He  uses  pi)  for  ov;  but  this  should  not  surprise  us,  as  he  was 
a  man  free  from  affectation  and  would  naturally  use  the  language 
as  it  was  spoken,  so  far  as  he  could  without  being  rude.2  But 
Lucian  is  not  fonder  of  the  prj  construction  in  questions  than  Dio 
Chrysostomus,  and  in  the  1301  pages  of  the  Teubner  edition  not  a 
single  example  of  apa  pi)  can  be  found,  prj  occurs  only  eight 
(really   seven)   times,   as    follows :   prj  dveipwv  viroKpnds  Tivas  i)pds 

vTre!.\r](f)€v;  (^Evvttvlov  I,  22  R.),  'AXXa  pr)  oveipos  /cai  ravra  eariv]  ("Oveipos 
II,  7°6)>  ov  6V  pr)  ku\  tov  2a>Kpa.Trjv  avTov  Ka\  tov  IlXdYeoi/a  etdes  iv  toIs 
veKpols',  (4?l\o\jsevbr)$  III,  52)>  'AXXa  pr)  'Eppa(pp68iTos  el]  ...  pr)  ovv  Ka\ 
o~v  toiovtov  tl  7T€Uov6as ',  ('EratpiKoi  Adyoi  III,  291))  prj  ti  tov  7rai8oTptj3rjv 
Aioripov  Xeyeiy;  (Ibid.  3°5)>    M  Tl  8ii)p.apr€S  fiaX&v  ',    (ty€vdoo~o(j)io-Tr)$    III, 

571),  and  one  in  the  Pseudo-Lucianic  dialogue  QiXoiraTpts  (III,  597), 

pr)  Tr)v   TeTpanTvv  (pfjS   Tr)v   UvOayopov ',    The    particles    peov,    apa   and    ap' 

ovv  are  found  occasionally. 

1  Dindorf  brackets  the  passage  in  which  apa  prj  with  the  subjunctive 
occurs  (XXVI,  524  R).  Dio  does  not  write  as  good  Attic  as  Niebuhr 
would  have  us  believe.     See  Amer.  Journ.  of  Philol.  I,  48,  50,  53,  57. 

2  See  A.  J.  P.  I,  47. 


434  /  E.   HARRY. 

Of  the  writers  of  the  third  century  A.  D.  I  selected  Plotinus 
and  Philostratus  for  investigation.  The  chief  representative 
of  Neo-Platonism  uses  apa,  apd  ye  and  ap'  ov,  but  never  pr)  or  dpa 
firj.  In  Philostratus  are  found  p5>v,  apa,  ap'  ov  and  7  ;  and  two  ex- 
amples of  pr)  I    Ap.  V,  33  prj  pel£6v  ti  Tovroiv]    V,  34  prj  ri  roiy  elprjpevoi? 

7rpo(TTidt]s;  In  the  thirty-ninth  epistle  another  question  (M^Se  ypdfaiv 
cpvydda  dve^;')  might  be  added  to  the  number. 

University  of  Cincinnati.  J«    ■£'•    rlARRY. 


9w 

by.  Z 


TKUBMOT 

OF  THE 

«WEM»TY  OP  ILUNOIS 

"NIVERSITY  OF  CINCINNATI 


Bulletin  No.  4 
October,   1901 


Publications  of  the  University  of  Qncinnati 

Series  II  Voi<.  I 


Observations  on  the  Efferent  Neurones  in  the  Electric 
Lobes  of  Torpedo  Occidentals  — 


SHINKISHI  Hatai 


The  University  Bulletins  are  Issued  Quarterly 


Entered  at  the  Post  Office  at  Cincinnati,  Ohio,  a^s  second-dass  matter 


UNIVERSITY  PRESS 
Cincinnati,  Ohio 


c 


tWuZ 

Neurones  in  Electric  Lobes  of  Torpedo. 


OBSERVATIONS  ON  THE  EFFERENT  NEURONES 
IN  THE  ELECTRIC  LOBES  OF  TORPEDO  OCCI- 
DENTALS. 

By  Shinkishi  HaTai, 
(From  the  Biological  Laboratory  of  the  University  of  Cincinnati.) 


Contents. 


I.    Materials    used    and    technique    employed    in    the    present 

investigations I 

II.     Finer  structure  of  the  efferent  neurones  of  the  electric  lobes 

in  Torpedo  occidentalis 2 

III.  Finer  structure  of  the  ground  substance  of  the  spinal  gan- 

glion cell  in  the  adult  white  rat 6 

IV.  Remarks  concerning  the  structure  of  the  ground  substance 

in  nerve  cells 10 

V.     Summary 11 

VI.     Illustrations 12 


I.    Materials  used  and  technique  employed 

IN   THE   PRESENT   INVESTIGATION. 

For  the  present  investigation,  the  efferent  neurones  in  the 
electric  lobes  of  Torpedo  occidentalism  and  the  spinal  ganglion 
cells  from  the  mid  -  cervical  ganglia  of  the  adult  white  rat 
were  used.  The  body  weight  of  the  rat  was  141  grams. 
The  torpedo  material,  which  was  generously  furnished  by 
Dr.  Ayers,  had  been  preserved  with  10%  formaline.  To  pre- 
pare this,  a  thin  piece  was  cut  from  the  lobe  and  transferred 
to  distilled  water  for  about  six  hours  in  order  to  remove  all 
the  formaline.  After  thorough  washing  with  water,  the 
material  was  transferred  to  35^  alcohol,  where  it  remained 
about  one  hour,  and  then  it  was  carried  through  graded 
alcohols  and  imbedded  in  paraffine  in  the  usual  way.  The 
sections  were  cut  12  ^  in  thickness.  For  staining,  a  satur- 
ated aqueous  solution  of  toluidin  blue,  and  for  contrast 
staining,  an  alcohol  solution  of  erythrosin,  were  used. 

Jour.  Cin.  Soc.  Nat.  Hist.,  Vol.  XX,  No.  1.       I  Printed  October  i,  1901. 


2  5.  Hatai. 

The  spinal  ganglion  of  the  white  rat  was  preserved  with 
the  author's  own  mixture  (formaline -acetic  sublimate  mix- 
ture) (*),  and  for  staining,  the  reagents  just  mentioned 
were  used. 

II.     FlNER  STRUCTURE   OF  THE  EFFERENT  NEURONES 

OF   THE   ELECTRIC   LOBES   IN   TORPEDO 

OCCIDENTALS. 

The  efferent  neurones  of  the  electric  lobes  of  Torpedo 
occidentalis  are  so  large,  more  than  o.i  mm.  in  diameter,  that 
they  can  easily  be  seen  with  the  naked  eye.  Under  moderate 
magnification,  the  cell  bodies  show  numerous  dendritic  pro- 
cesses and  the  single  axone  is  also  visible  in  most  cases. 

The  general  form  of  the  cell  body  is  somewhat  similar  to 
that  of  the  motor  cells  in  the  ventral  horn  of  the  spinal  cord 
in  man  and  the  higher  mammals.  In  most  cases,  the  nucleus 
lies  on  the  side  of  the  cell -body  towards  the  axis- cylinder 
process.  The  nucleus  is  nearly  spherical,  and  very  large  in 
size  proportionately  to  the  cell-body  (40-30  (*).  The  arrange- 
ment of  the  chromosomes  in  the  nucleus  is  somewhat 
peculiar.  They  do  not  show  minute  spherules  suspended  in 
the  delicate  meshwork  of  the  linin  substance,  but  instead  of 
that,  irregular  large  masses  which  fill  up  meshes  of  the  linin. 

The  nucleolus  is  always  visible  and  lies  at  one  pole  of 
the  nucleus.  Curiously  enough,  the  nucleolus,  as  a  rule,  lies 
in  the  same  relative  position  in  all  the  cells  of  a  given 
section. 

Under  the  higher  magnification,  the  internal  structure  of 
the  cell-body  shows  a  fibrillar  arrangement  of  the  cytoplasm. 
The  nature  of  this  fibrillar  structure  will  be  discussed  later 
on.  In  this  chapter,  only  the  general  arrangements  of  these 
fibrils  will  be  described. 

Briefly  speaking,  the  cell-body,  except  the  nucleus  presents 
everywhere  a  fibrillar  arrangement  of  the  cytoplasm.  The 
following  descriptions  apply  to  the  serial  sections  of  one  cell 
(102  [i  in  diameter,  and  60  p.  in  thickness),  and  give  a 
general  idea  of  the  structure  above  mentioned. 


(*)  Hatai,  S. —  Finer  structure  of  the  spinal  ganglion  cells  in  the  white  rat. — Jour, 
of  Comp.  Neurology,  Vol.  XI,  No.  I,  1901. 


Neurones  in  Electric  Lobes  of  Torpedo.  3 

Fig.  1  is  a  section  passing  through  the  periphery  of  the 
cell-body.  In  this  figure,  the  dendritic  processes  are  shown, 
but  not  the  neuraxone.  The  position  where  the  neuraxone 
will  arise  in  the  sections  is  marked  by  A.  The  fibrillar  bun- 
dles which  come  from  all  dendritic  processes  of  one  side  of 
the  cell-body  (a)  take  a  curving  course  toward  the  axone 
hillock,  thus  forming  an  arrangement  like  an  inverted  U. 
Other  fibrillar  bundles  come  also  from  the  dendrites  on  the 
other  side  (b)  and  take  the  same  course  toward  the  neuraxone. 
The  dotted  areas  are  interpreted  as  the  cross -sections  of 
the  similar  fibrillar  bundles  which,  running  through  the  cell- 
body  in  different  directions,  are  therefore  cut  at  different 
angles.  In  this  figure,  the  fibrillar  bundles  connecting  the 
dendrites  with  each  other  are  shown  very  poorly. 

Fig.  2  is  the  section  nearer  the  center  of  the  cell-body  and 
follows  Fig.  1.  In  this  figure,  the  four  dendrites  are  shown 
clearly,  and  the  localities  of  the  neuraxone  is  indicated  by 
"A,"  although  it  does  not  appear  at  this  level.  The  fibrillar 
bundles  which  form  the  neuraxone  come  from  each  of  the 
dendrites.  The  dendrites  themselves  have  close  relations 
with  each  other  by  means  of  the  connecting  fibrillar  bundles 
passing  between  them.  The  nucleus  is  surrounded  by  the 
fibrils  coming  from  one  of  the  dendrites  (V).  The  fibrillar 
bundles  which  come  from  the  dendrites  (d)  also  take  a  part 
in  investing  the  nucleus.  The  cross-sections  of  the  fibrillar 
bundles  show  as  clearly  separated  groups. 

Fig.  3  is  a  section  passing  through  the  middle  of  the 
nucleus  and  follows  Fig.  2.  In  this  figure,  the  nucleolus  is 
visible.  The  fibrillar  arrangements  are  slightly  different 
from  those  in  the  figures  already  given.  In  this  section  the 
fibrils  do  not  form  large  bundles,  but  are  divided  into  smaller 
strands  and  interwoven.  The  intimate  connections  between 
the  dendrites  are  clearly  shown.  The  nucleus  is  also  sur- 
rounded by  the  bundles  of  the  fibrils,  which  come  from  some 
of  the  dendrites.  As  a  rule,  in  this  level  the  fibrillar  bundles 
near  the  nucleus  are  short,  because  bundles  are,  for  the  most 
part,  cut  more  or  less  at  right  angles  to  their  long  axis.  This 
suggests  that  the  fibrillar  bundles  converge  towards  the 
nucleus.     The  peculiar  arrangement  of  the  fibrils  near  the 


4  S.  Hatai. 

nucleus  has  been  described  as  "  vortex "  or  "  spiral,"  or 
sometimes  "  Gitterahnliche  Anordnung."  On  the  contrary, 
the  fibrillar  bundles  at  the  periphery  present  comparatively 
long  sections.  In  this  section,  the  neuraxone  is  not  yet 
shown. 

Fig.  4  is  a  section  of  the  cell  -  body  at  another  level.  In 
this  figure,  three  dendrites,  nucleus,  and  neuraxone  are 
clearly  shown.  The  neuraxone  "A"  lies  at  one  corner  of  the 
base  of  the  rectangular  cell-body.  An  intimate  connection 
of  each  dendrite  with  that  of  the  other,  and  also  of  all  the 
dendrites  with  neuraxone'  is  clearly  shown  in  this  figure. 
A  curious  arrangement  of  the  fibrils  is  noticeable  very  near 
the  axone  hillock,  where  the  fibrillar  bundles  have  a  beauti- 
ful spiral  arrangement.  This  spiral  arrangement  is  produced 
by  the  fibrils  coming  from  various  dendrites  as  is  shown  in 
the  illustration.  In  this  figure,  connecting  fibrils  between 
the  dendrites  (a)  and  (c)  are  shown  very  clearly. 

Fig.  5  is  a  section  passing  through  the  periphery  of  the 
side  opposite  to  that  shown  in  Fig.  i.  In  this  figure,  four 
dendritic  processes  are  plainly  shown  —  one  from  each  corner 
of  a  somewhat  rectangular -shaped  cell-bod}^.  The  position 
from  where  the  neuraxone  will  arise  in  other  section  is 
marked  by  "A."  A  clear  oblong  space  near  the  center  of 
the  cell-body  is  the  place  where  the  nucleus  lies  in  the  other 
sections.  The  fibrillar  bundles  which  come  from  the  den- 
drite (a)  run  towards  the  dendrites  (c,  d)  along  the  one  side 
of  the  nucleus,  and  finally  enter  the  dendrites  (c,  d).  Along 
the  course,  a  few  small  fibrillar  bundles  diverge  towards  the 
periphery  of  the  cell -body.  The  fibrillar  bundles  which 
come  from  the  dendrites  (b)  run  toward  the  dendrites  (c,  d~) 
in  a  somewhat  similar  manner  to  those  from  the  dendrite  (a). 
In  this  case,  the  fibrillar  bundles  divide  into  two  branches  at 
the  nucleus  and  after  encircling  the  nucleus,  they  enter  in  the 
dendrites  (c,  d)  and  become  continuous  with  those  from  the 
dendrite  (a).  From  the  base  of  the  dendrite  (&),  small  fibrillar 
bundles  are  distributed  toward  the  neuraxone.  From  the 
dendrites  c,  d,  the  bundles  of  fibrils  arise,  and  run  toward  the 
neuraxone.  Along  their  course,  these  bundles  are  increased 
by  the  addition  of  numerous  bundles  of  fibrils  which  come 


Neurones  in  Electric  Lobes  of  Torpedo.  5 

from  the  periphery  of  the  cell-body  to  form  the  yet  larger 
bundles  found  in  the  axone  hillock.  The  dendrites  a  and  b 
are  subdivided  into  two  branches.  In  this  case  the  branches 
are  also  connected  by  a  few  fibrils.  These  branches  which 
are  divided  from  the  main  dendrites  {a,  b})  receive  fibrils  from 
various  regions  of  the  cell-body. 

From  the  above  description,  two  important  relations  are 
evident :  (1)  That  each  dendrite  is  connected  by  the  fibrillar 
bundles  with  several  and  possibly  all  the  others,  and  (2)  in 
each  case,  the  nucleus  is  partially  surrounded  or  encircled  by 
the  fibrillar  bundles,  on  their  way  from  the  dendrites  to  enter 
into  the  neuraxone. 

As  a  rule,  the  fibrils  in  the  dendrites  are  very  conspicuous, 
presenting  long  continuous  lines,  while  in  the  cell-body  they 
take  tortuous  or  irregular  courses,  so  that  the  cross-section 
of  the  cell-body  presents  minutely  dotted  areas,  representing 
the  cross-section  of  the  bundles.  From  this,  it  is  inferred 
that  the  entire  course  of  some  of  the  bundles  must  be  very 
complex. 

Fig.  6  is  a  diagram  reconstructed  from  the  serial  sections 
of  the  cell-body  in  order  to  depict  schematically  its  structure 
and  to  show  the  fibrillar  tracts  distributed  throughout  it. 
L,et  us  take  any  one  of  the  dendrites  from  the  Fig.  6,  and 
trace  the  lines  which  represent  the  fibrillar  bundles.  In  the 
dendrite  (d),  black  continuous  lines  present  the  out-going 
fibrillar  bundles,  while  dotted  lines  in  the  same  dendrite 
represent  the  in-coming  fibrillar  bundles  from  other  dendrites. 
If  we  trace  one  of  the  black  lines  (3),  it  enters  into  the  den- 
drites which  lie  in  both  sides,  and  other  black  lines  (1)  run 
toward  the  nucleus  and  partially  encircle  it.  The  fibrils  con- 
tinue from  the  nucleus  toward  the  axone  and  finally  enter 
into  the  axis  cylinder.  In  the  remaining  dendrites,  the 
fibrillar  tracts  are  just  the  same  in  their  distribution  with 
those  of  dendrites  (d). 

In  some  cases,  the  fibrillar  bundles  which  run  from  the 
dendrite  not  only  enter  into  the  dendrites  which  lie  nearest 
on  both  sides,  but  they  also  connect  with  other  dendrites 
further  distant  (2).  In  the  cross-section  of  the  cell-body,  we 
notice  very  often  the  following  appearance :     The  neighbor- 


6  S.  Hatai. 

hood  of  the  nucleus  is  composed  of  peculiarly  arranged 
fibrils,  forming  a  " spiral"  or  "swirl."  These  appearances 
are  caused  by  the  fibrils,  which  take  very  irregular  courses 
and  partially  encircle  the  nucleus  in  a  tortuous  manner. 

III.     FlNER  STRUCTURE  OF  THE  GROUND  SUBSTANCE 

OF  THE  SPINAS  GANGLION  CELLS  IN  THE 

ADULT  WHITE   RAT. 

It  remains  to  discuss  the  real  nature  of  the  fibrillar  struc- 
tures mentioned  above,  and  to  this  end  the  structure  of  the 
ground  substance  of  the  nerve-cells  must  first  be  considered. 

Concerning  the  structure  of  the  ground  substance  in  nerve- 
cells,  two  main  views  are  held:  the  "fibrillar"  and  "  non- 
fibrillar"  structure.  The  former  theory  may  also  be  sub- 
divided. One  view  is  represented  by  the  theory  of  Bethe  (*) 
who  regards  the  ground  substance  as  composed  of  "  Peri 
Fibrillar  Substanz"  and  "Fibrillen."  The  so-called  Fibrillen 
are  independent  individuals  distributed  throughout  the  cell- 
body  in  a  certain  way,  where  they  neither  anastomose  nor 
branch.  Another  fibrillar  theory  is  that  of  Apathy  (1). 
According  to  this  author,  the  primitive  neurofibrils  are  to  be 
distinguished  by  means  of  special  technique,  in  the  nerve- 
cells  as  Bethe  describes.  These  fibrils  however,  are  not 
isolated,  but  are  connected  with  each  other  by  means  of  deli- 
cate branches,  thus  forming  a  very  complicated  anastomosis 
within  the  nerve-cells. 

The  non-fibrillar  theories  may  also  be  divided  into  two 
groups,  represented  by  the  theory  of  Apathy  (1),  Nansen 
(2),  Butschli  (3),  etc.  Nansen  holds  the  view  of  primitive 
tubular  structure  of  the  formation  of  the  ground  substance 
of  the  nerve-cells,  that  is,  the  ground  substance  is  entirely 
composed  of  extremely  small  tubules  which  are  directly 
continuous  with  the  neuraxone. 


(*)  Bethe,  A.— Uber  die  Primitiv  Fibrillen  in  den  Ganglien-zellen  von  Menschen 
und  Wirbelthieren. —  Arch,  fur  Mikrosk.  Anat.,  Bd.  51. 

(1)  Apathy. —  Das  leitende  Element  des  Nervensystems,   u.  s.  w.  —  Mitheil.  d. 
Zoolog.  Station  zu  Neapel,  B'd  XII,  '97. 

( 2 )  Nansen,  F.— The  structure  and  combination  of  the  histological  elements  of 
the  central  nervous  system. —  Bergeu,  '87. 

(3)  Butschli.—  Investigations  on    microscopic    forms  and  on  protoplasm.— '94. 
Translation  to  English. 


Neurones  in  Electric  Lobes  of  Torpedo.  7 

Butschli,  Held  (x),  Van  Gehuchten  (2),  Von  Lenhossek  (s), 
Ramon  y  Cajal  (4),  Marinisco  (5),  Ewing  (6),  a.  o.,  hold 
the  view  of  reticular  or  spongy  formation  of  the  ground 
substance,  stating  that  the  fibrillar  structure  described  by 
others  are  not  true  fibrils  but  rows  of  fine  granules  which 
form  the  reticular  arrangement  of  the  ground  substance. 

The  writer's  observations  on  this  subject  are  as  follows: 
The  ground  substance  of  the  spinal  ganglion  cells  of  the 
white  rat  exhibits  a  reticular  structure  as  shown  in  Fig.  7. 
The  meshes  of  the  reticulum  are  very  small  but  conspicuous. 
The  size  and  form  of  the  meshes  vary.  Generally,  in  the 
clear  zone  at  the  periphery  of  the  cell-body,  the  meshes  are 
always  larger  and  more  conspicuous  than  in  the  remaining 
part.  In  the  neighborhood  of  the  axone  hillock  the  meshes 
are  not  only  much  diminished  in  size,  but  also  they  are  much 
elongated  along  one  axis.  Around  the  nucleus,  the  meshes 
reach  a  minimum  size.  The  form  of  the  reticulum  at  the 
periphery  shows  meshes  of  a  somewhat  polygonal  shape,  but 
in  the  remaining  part  of  the  cell  these  meshes  are  elongated, 
especially  around  the  nucleus  and  near  the  neuraxone.  Upon 
examining  with  a  higher  magnification,  the  protoplasmic 
threads  or  filaments  which  forms  the  reticulum,  we  see  that 
it  is  not  smooth  but  has  a  somewhat  varicose  appearance,  due 
to  the  presence  of  small  bead -like  arrangements  on  the 
course  of  the  filaments.  This  bead  was  called  by  Held  (*) 
a  "  neurosome,"  who  discovered  the  occurrence  of  the  neuro- 
some  not  only  at  the  connecting  point  of  the  net  but  also 
inside  the  net.  The  writer  noticed  the  occurrence  of  these 
structures  not  only  at  the  connecting  points  of  the  net  but 
also  in  the  course  of  the  filament,  but  could  not  find  them 
inside  the  reticulum. 


(1)  Held. —  Beitrage  zur  Strukturen    der    Nerven-zellen  und  ihren   Fortsatze. — 
Erste   Abhandlung.     Arch,  fur  Anat.  und  Entwickelungs.      Anat.  Abth.,  '95. 

(2)  Van  Gehuchten. —  Anatomie  du  system  nerveux  de  l'homme. —  I,auvain,  1894. 

(3)  Von  I^enhossek. —  Feinere  Bau  des  Nervensystems.— '95.    P.  147. 

(4)  Cajal. —  Fstructura    del    protoplasma    nerviso.  —  Revista    trimestral    micro- 
grafica,  Vol.  I,  fasc.  1,  '96. 

(5)  Marinisco.—  Pathologie  generate  de  la  cellule  nerveuse.—  I,a  Presse  M6di- 
cale,  '97. 

(6)  Ewing.—  Studies  on  ganglion  cells.— Arch,   of  Neurol,  and   Psychopathol. , 
Vol.  I,  No.  3.     '98. 

(*)  Held.—  I,oc.  cit. 


8  S.  Hatai. 

This  bead  or  neurosome  has  peculiar  chemical  affinities  for 
the  staining  fluids.  Bosin  or  erythrosin  stain  this  element 
very  deeply,  so  that  it  can  easily  be  distinguished  from  the 
rest  of  structures.  The  fine  filament  joining  these  beads 
seems  to  be  slightly  different  from  the  neurosome  itself,  as  is 
shown  by  a  slightly  different  staining  reaction.  It  seems, 
indeed,  that  these  neurosomes  are  a  highly  differentiated 
portion  of  the  protoplasm  which  forms  the  reticulum. 

The  form  and  size  of  the  neurosomes  are  different  in 
different  localities,  as  has  been  already  described  by  Held. 
These  structures  are  especially  numerous  within  the  axone 
hillock  and  intracellular  extension  of  the  axone.  At  the 
periphery  of  the  spinal  ganglion  cells,  the  individual  meshes 
of  the  reticulum  are  so  large  that  the  neurosomes  are  less 
crowded,  hence,  in  this  region,  they  are  scattered  very  irregu- 
larly. But  on  the  contrary,  in  the  remaining  parts  of  the 
cell,  the  meshes  of  the  reticulum  are  elongated  in  shape  and 
the  rows  of  neurosomes  become  more  crowded  together,  thus 
giving  the  fibrillar  appearence.  At  first  glance,  this  arrange- 
ment of  neurosomes  looks  very  much  like  the  fibrils  which 
have  been  described  by  many  authors.  Careful  observations, 
however,  show  that  these  lines  appearing  like  fibrils  are 
composed  of  a  row  of  minute  beads  arranged  serially. 
Moreover,  these  pseudo- fibrils  are  connected  by  protoplasmic 
threads,  thus  forming  the  reticulum.  This  structure  is  shown 
in  Fig.  7.  Around  the  nucleus  these  neurosomes  form  some- 
what concentric  lines  in  a  very  beautiful  manner.  But 
gradually  the  figure  becomes  irregular  as  the  reticulum 
approaches  the  periphery.  This  is  the  appearance  generally 
found  in  the  spinal  ganglion  cells.  Sometimes  the  cell  shows 
different  arrangement  of  neurosomes,  namely,  concentric 
lines  at  the  periphery  but  not  in  the  neighborhood  of  nucleus. 
Still  other  variations  in  arrangement  are  found. 

Graf  (*)  noticed  the  fibrils  which  are  composed  of  a  row  of 
minute  beads,  in  the  Purkinji  cells  of  human  cerebellar  cor- 
tex. He  said:  "  The  cytoplasma  show  the  most  beautiful 
fibrillar  structure  that  I  have  ever  seen.     The  fibrillse  are 


(*)  Graf,  A.— On  the  use  and  properties  of  a  new  fixing  fluid  (chrom  -oxalic.) 
Bull,  of  Pathol.  Institute  of  the  New  York  Hospitals,  '97.  Vol.  II,  p.  386. 


Neurones  in  Electric  Lobes  of  Torpedo.  9 

exceedingly  fine  and  are  very  regularly  arranged  in  the  cell- 
processes  and  on  the  surface  of  the  cell,  whereas  they  form  a 
more  intricate  network  in  the  center  of  the  cell,  especially 
around  the  nucleus.  By  closer  observation  of  a  favorable  spot 
(the  best  places  are  where  the  stain  is  not  very  intensive) 
we  notice  that  the  finest  cytoplasmic  fibrillse  are  not  smooth, 
like  smooth  muscle  fibrils,  for  instance,  but  are  composed  of 
a  row  of  minute  beads  closely  arranged  in  single  file." 

Held  believes  that  the  fibrils,  according  to  some  investi- 
gators, are  in  reality  identical  with  rows  of  neurosomes.  He 
hints  that  some  of  the  fibrils  represent  bands  of  neurosomes; 
other  fibrils  described  by  Flemming  are  bundles  of  cyto- 
spongium. 

My  own  observations  support  Held's  suggestion.  My  prep- 
arations show  sometimes  exactly  the  fibrillar  structure  de- 
scribed by  Graf,  and  I  find  this  condition  in  the  efferent 
neurones  of  the  Torpedo,  as  well  as  in  the  spinal  ganglion 
cells  in  the  white  rat.  These  fibrils  can  always  be  resolved 
into  rows  of  neurosomes. 

Another  important  point  is,  that  the  meshes  of  the  reticu- 
lum in  the  cell-body  become  more  and  more  elongated  toward 
the  axis  cylinder.  Thus  it  looks  as  if  the  fibrils  are  radiating 
from  the  axone  around  the  nucleus. 

The  peculiar  character  of  the  region  from  where  the  axis 
cylinder  originates  was  first  described  by  Schaffer  (1). 

This  region  of  the  cell-body  he  called  the  "axone  hillock." 
It  is  admitted  by  most  investigators  that  the  axone  hillock, 
as  well  as  the  axis  cylinder,  show  a  parallel  arrangement  of 
cytoplasm.  The  writer  notices  also  these  arrangements  of  fine 
cytoplasmic  threads,  which  carry  the  neurosomes,  showing  a 
convergent  arrangement  toward  the  axis  cylinder.  In  this 
region  the  meshes  of  the  reticulum  are  very  small,  but  care- 
ful examination  shows  that  the  axone  hillock,  as  well  as  axis 
cylinder,  are  composed  of  an  altered  reticulum. 

The  arrangement  of  neurosomes,  except  in  the  axone  hil- 
lock, is  not  the  same  in  all  nerve-cells,  but  differs  according 
to  the  type  of  the  cells. 


(1)  Schaffer,  K. —  Kurze  Anmerkung  iiher  die  Morphologische  Differenz  des  Axen 
Cylinders  in  Verhaltnisse  zu  dem  Protoplasmatischen  Fortsatze  bei  Nissl's 
Farbung. —  Neurol.  Centralbl.,  Leipzig,  Bd.  XII,  '93,  S.  849-851. 


io  5.  Hatai. 

In  the  motor  ganglion  cells  in  the  anterior  horn  of  the 
spinal  cord,  the  neurosome  presents  quite  a  different  arrange- 
ment from  that  of  spinal  ganglion  cells.  In  the  former  group 
the  meshes  of  the  reticulum  do  not  show  the  honey-comb 
form,  but  an  elongated  shape.  The  cytoplasmic  thread  car- 
ries a  great  number  of  neurosomes,  which  form  straight 
chains.  These  chains  run  parallel  to  the  periphery  toward 
the  dendrites,  as  well  as  toward  the  axis  cylinder.  Around 
the  nucleus,  however,  these  chains  have  the  arrangement 
found  in  the  spinal  ganglion  cells. 

The  Purkinjii  cells  in  cerebellar  cortex  in  the  white  rat 
show  still  a  different  arrangement  of  neurosomes.  In  these 
cells  the  neurosomes  accumulate  at  the  base  of  the  main  den- 
drites, showing  very  intricate  arrangement.  But  near  the 
entrance  of  the  dendrites  the  irregular  chains  rearrange 
themselves,  forming  a  regular  line  of  neurosomic  fibrils.  The 
remaining  part  of  the  cell-body  show  nearly  the  same  arrange- 
ment as  that  of  the  spinal  ganglion  cells. 

IV. — Remarks  concerning  the  structure  of  the 

GROUND  SUBSTANCE  IN  NERVE  CELLS. 

As  has  been  mentioned  already,  the  ground  substance  of 
the  spinal  ganglion  cells  of  the  white  rat  presents  very  clearly 
the  reticular  structure.  This  structure,  however,  is  altered 
by  the  growth  of  cell-body;  for  example,  the  prolongation  of 
the  axis  cylinder  from  the  cell-body  is  accompanied  by  an 
elongation  of  the  primitively  polygonal  meshes  of  the  reticu- 
lum, thus  giving  a  fibrillar  appearance  to  the  ground  sub- 
stance. 

The  same  holds  true  in  the  case  of  the  Torpedo.  The  ap- 
parent fibrils  result  from  alterations  in  the  reticulum,  and, 
therefore,  should  not  be  compared  to  those  of  Bethe's.  Al- 
though, in  the  case  of  the  Torpedo,  the  reticulum  is  hard  to 
see,  yet  it  is  sometimes  clearly  demonstrable  in  thin  sections 
properly  stained. 

In  the  spinal  ganglion  cells  of  the  higher  mammalia,  except 
in  Dogiel's  second  type  of  cells,  the  cell -body  sends  off  only 
one  prolongation,  while  in  the  case  of  Torpedo,  the  efferent 

io 


Neurones  in  Electric  Lobes  of  Torpedo.  n 

neurones  of  the  electric  organ  give  numerous  processes  from 
the  cell-body.  In  the  former  case,  the  meshes  of  the  reticu- 
lum are  changed  gradually  from  a  regular  polygonal  form 
to  those  much  drawn  -  out  in  the  axone  hillock.  In  the 
case  of  the  Torpedo,  however,  the  arrangement  of  the 
reticulum  is  modified  not  only  toward  the  axis  cylinder,  but 
in  every  part  of  the  cell-body  from  which  dendritic  processes 
arise.  The  appearances  in  Torpedo  can  be  explained  as  a 
result  of  the  growth  changes  of  the  cell-body.  Judging  from 
what  we  find  in  the  rat,  we  assume  in  the  first  place  the  spinal 
ganglion  cell  to  be  a  spherical  mass  filled  by  the  wide  meshed 
reticulum.  For  the  same  reason  we  assume  that  this  spheri- 
cal mass  is  pulled  out  at  each  point  where  there  is  a  dendrite, 
and  thus  modified  as  it  is  where  the  neuraxone  is  formed 
from  the  axone  hillock.  As  a  result,  the  primitive  polygonal 
meshes  are  transformed  mechanically  by  the  growth  changes 
and  thus  give  rise  to  the  fibrillar  appearance.  If  numerous 
processes  are  formed  by  the  cell,  as  in  the  case  of  Torpedo, 
then  the  resulting  appearance  is  quite  complex.  But  the 
principle  of  its  formation  is  the  same  as  in  the  more  simple 
spinal  ganglion  cell.  The  so-called  fibrillar  arrangement 
in  the  writer's  preparation  is  thus  explained: 

V. — Summary. 

i.  The  efferent  neurones  of  the  electric  lobes  of  Torpedo 
occidentalis  present  a  fibrillar  appearance  of  the  ground 
substance. 

2.  This  appearance,  however,  is  due  to  an  alteration  in  the 
shape  of  the  meshes  of  the  reticulum,  and,  therefore,  it  can- 
not be  compared  with  the  fibrils  described  by  Bethe,  Apathy, 
and  others. 

3.  The  meshes  of  the  reticulum,  which  are  regarded  as  the 
primitive  by  the  present  writer,  are  altered  by  the  growth  of 
the  cell-body  where  the  processes,  both  axone  and  dendrite, 
arise  and  become  extremely  elongated  in  these  branches. 

4.  Gradations  from  the  primitive  shape  of  the  meshes  to 
the  altered  form  which  appears  fibrillar,  are  clearly  visible  in 
the  spinal  ganglion  cells  of  the  white  rat. 

11 


5.  Hatai. 


VI. — Illustrations.     (Plate  I.) 

Fig.  1-5 — Five  serial  sections  from  a  single  efferent  neurone  in  electric 
lobe  of  Torpedo  occidentalis.  Mean  diameter  of  the  cell- 
body  (120  p.  x  83  p)  ;   of  the  nuclei  (37  p  x  34  p). 

Fig.  6 — Diagram   showing  the   fibrillar  arrangement   of  the   efferent 
neurone  in  an  electric  lobe  of  Torpedo  occidentalis. 

Fig.  7 — Spinal  ganglion  cell  from  the  mid-cervical  ganglia  of  the  adult 
white  rat.     Cell-body  (41  p  x  30  p.) ;  nucleus  (15  p  x  15  p). 


12 


University  of  Cincinnati 

Buixetin  No.  4.     October,  1901. 


Plate  1 


Hatai  Efferent  Neurones. 


LIBRARY 


ILLliHUr, 


